Copyright © 2000 by The Resilience Alliance.

Sutherland, G. D., A. S. Harestad, K. Price, and K. P. Lertzman. 2000. Scaling of natal dispersal distances
in terrestrial birds and mammals. Conservation Ecology 4(1): 16. [online] URL:
http://www.consecol.org/vol4/iss1/art16

Report
Scaling of Natal Dispersal Distances in Terrestrial Birds and
Mammals
Glenn D. Sutherland1, Alton S. Harestad2, Karen Price2, and Kenneth P. Lertzman2

ABSTRACT. Natal dispersal is a process that is critical in the spatial dynamics of populations, including
population spread, recolonization, and gene flow. It is a central focus of conservation issues for many vertebrate
species. Using data for 77 bird and 68 mammal species, we tested whether median and maximum natal dispersal
distances were correlated with body mass, diet type, social system, taxonomic family, and migratory status. Body
mass and diet type were found to predict both median and maximum natal dispersal distances in mammals: large
species dispersed farther than small ones, and carnivorous species dispersed farther than herbivores and
omnivores. Similar relationships occurred for carnivorous bird species, but not for herbivorous or omnivorous
ones. Natal dispersal distances in birds or mammals were not significantly related to broad categories of social
systems. Only in birds were factors such as taxonomic relatedness and migratory status correlated with natal
dispersal, and then only for maximum distances. Summary properties of dispersal processes appeared to be
derived from interactions among behavioral and morphological characteristics of species and from their linkages
to the dynamics of resource availability in landscapes.

In all the species we examined, most dispersers moved relatively short distances, and long-distance dispersal was
uncommon. On the basis of these findings, we fit an empirical model based on the negative exponential
distribution for calculating minimum probabilities that animals disperse particular distances from their natal areas.
This model, coupled with knowledge of a species' body mass and diet type, can be used to conservatively predict
dispersal distances for different species and examine possible consequences of large-scale habitat alterations on
connectedness between populations. Taken together, our results can provide managers with the means to identify
species vulnerable to landscape-level habitat changes such as forest fragmentation. In addition, our dispersal
models can be used to predict which species in a community are likely to be the most vulnerable to loss of
connectedness and allow managers to test the merits of alternative habitat conservation plans.

INTRODUCTION dispersal occurs regularly, but at a relatively low

Distances moved by juvenile animals during natal
dispersal are a fundamental element of demography
(Arcese 1989), population dispersion, colonization
(Hengeveld 1994), and gene flow (Neigel and Avise
1993, Nelson 1993). Despite their ecological
importance, dispersal movements are among the least
understood attributes of both individual animals and
populations. Natal dispersal distances vary
considerably among species (Swingland 1982). The
frequency of dispersal decreases with increasing
distance from the natal area (Taylor 1980, Paradis et
al. 1998). Available evidence indicates that short
dispersal distances are frequent and strongly influence
age and sex structure, abundance, and relatedness
within populations. In many species, long-distance
frequency. Nonetheless, long dispersal distances are
important in invasion and recolonization processes
(Shaw 1995) and in the genetic structuring of
populations (Ibrahim et al. 1996).
No consensus exists on the factors that determine the
distribution of dispersal distances moved by juvenile
animals (Paradis et al. 1998). Dominant hypotheses
currently used to explain the ultimate benefits of natal
dispersal are intrasexual competition for resources
(e.g., mates, food, and space) and inbreeding
avoidance (Dobson 1982, Pusey 1987, Wolff 1993).
However, there is considerable controversy about the
relative roles of these two processes in structuring
dispersal patterns within and among species
(Bengtsson 1978, Moore and Ali 1984, Dobson and

1
University of British Columbia; 2Simon Fraser University

Jones 1985, Shields 1987, Wolff 1993, Kunkele and
von Holst 1996). All hypotheses assume that
dispersers incur costs to survival and/or fecundity
(Bengtsson 1978) and that these costs increase with
distance traveled. Dispersing individuals may face
increased mortality risks associated with unfamiliar
habitats, passage through areas of relatively high
predator densities, or the physiological costs of
extensive movement (Waser et al. 1994, Plissner and
Gowaty 1996).
Lack of quantitative methods for predicting the spatial
scale of dispersal from natal habitats is a major
limitation in the development of theories and tools for
forecasting the effects of landscape alteration on
connectivity between habitats and subpopulations of
animals. Recent spatially explicit models for assessing
the effects of habitat fragmentation on populations of
animals require knowledge of the dispersal
characteristics of their target species. For example, two
key parameters are the dispersal rate (number of
dispersers leaving their natal territory) and the
expected distance dispersed by each disperser (Pulliam
et al. 1992, With and Crist 1995, Schumaker 1996).
However, dispersal characteristics of this type are
known in detail for only a few species. While simple
mathematical models have been fit to the dispersal
distributions of some of these species (Waser 1987,
Miller and Carroll 1989, Caley 1991), generalizing
them to other species is questionable. Parameter
estimates vary widely among species (Miller and
Carroll 1989), and many models assume particular
behavioral mechanisms governing the distance moved
by individual dispersers that may themselves not be
general (Rees 1993). Interstudy differences in the
intensity of sampling and the high probability of biases
introduced through restricted searching patterns
(Porter and Dooley 1993) further reduce the generality
of these models.
The prominent role of dispersal in the life history of
most species suggests that relationships exist between
dispersal patterns and basic life-history attributes of
species. Indeed, some have recently been described:
mean and standard deviation (SD) of natal and
breeding dispersal distances in many species of British
birds depend on habitat type and migratory capability,
with body size as a covariate (Paradis et al. 1998).
However, for some species of mammals, median
dispersal distance appears to be related to diet type and
body size (Van Vuren 1998). We wished to place the
analysis of these relationships into a common
ecological framework and determine general,
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predictive relationships for birds and mammals.
Accordingly, we developed a number of a priori
expectations regarding how attributes of species should
relate to their ability to disperse a given distance:
1. Dispersal distance should increase with
increasing body size. Scaling functions that
relate body size to interspecific variation in
biological functions ranging from rates of
cellular metabolism to population dynamics are
common (Peters 1983, Schmidt-Nielsen 1984,
Holling 1992, Silva 1998). Generally, such
functions can be explained in terms of how
individuals acquire and use resources as a
function of body size (Brown and Maurer
1989, Kozlowski and Weiner 1997, West et al.
1997). In particular, allometric scaling
equations with an exponent value of
approximately ¾ are expected for size-
dependent resource utilization relationships
(McNab 1963, Peters 1983, West et al. 1997).
The dominant hypotheses about the ultimate
benefits of dispersal can also be interpreted
broadly in terms of the availability of resources
(ecological or genetic). Larger animals also
have more time, on average, to explore (both
between meals and over a lifetime) as longevity
increases with body mass ( Peters 1983). Both
explanations lead to the prediction that larger
species will disperse farther (assuming some
benefit to dispersing); thus, across species we
expected body mass to be significantly
correlated with dispersal distance.
2. Dispersal distance should increase with
increasingly exclusive acquisition of resources.
We examined two correlates of spacing
behavior: diet type and social system. Diets
vary in resource abundance and predictability,
and diet type influences spacing behavior
(Harestad and Bunnell 1979). Because
utilizable energy per unit area is greater at
lower trophic levels (Harestad and Bunnell
1979) and because home range size increases
with decreasing resource abundance (Mace et
al. 1982), we predicted that predators would
disperse farther than omnivores and that
omnivores would disperse farther than
herbivores. In both birds and mammals, factors
such as type of mating system and form of
territorial defense can influence the relative
distance moved by dispersers (Greenwood
1980). For example, if males defend resources,
females usually disperse farther (Greenwood

and Harvey 1982). If both sexes defend
territories, gender bias in dispersal does not
occur in some species (Mattysen and Schmidt
1987, Arcese 1989). Males are likely to
disperse farther in polyandrous species (Oring
and Lank 1984) and if males defend females
(Lessels 1985). We predicted that species
defending breeding territories should disperse
farther than species with overlapping or non-
exclusive territories. Patterns of dispersal
distances in more gregarious species (e.g.,
colonial nesters) could potentially differ
substantially from those of nongregarious
species, depending on the density, geographic
range, and aggregation patterns of breeding
sites within and among colonies. For example,
median distances may be quite short if colonies
are clumped in distribution, but maximum
distances may be very long if colonies are
sparsely dispersed at a large (e.g., continental)
geographic scale.
3. In birds, dispersal distances for migrants
should exceed those for residents. Recent
evidence suggests that the breeding dispersal
distances for migrant species are longer than
those for resident species (Paradis et al. 1998),
although this result was not found for natal
dispersal distances.
4. Dispersal distances for closely related species
should be more similar than for less closely
related species. Interspecific comparisons of
ecological relationships are often confounded
because species are part of a hierarchically
structured phylogeny and cannot be regarded,
for statistical purposes, as if drawn
independently from the same distribution
(Felsenstein 1985). In our study, evolutionary
factors determining body size, although likely
independent of dispersal, may be strongly
correlated in closely related species.
If the dispersal distance varies predictably with these
life-history attributes, then researchers could use such
relationships to identify species potentially vulnerable
to loss of connectivity between habitats, and habitat
planners could use these relationships to assess the
potential risks of alternative habitat configurations to
locally vulnerable species.
In this paper, we define dispersal as the movement of
an individual out of an area larger than its home range,
with no predictable returns, i.e., excluding migrations
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(Bunnell and Harestad 1983). For most species and
most dispersing individuals, dispersal takes place
before first reproduction and is termed natal dispersal
(Howard 1960). Natal dispersal is usually the single
largest (and often only) long-distance movement made
by individual animals (Dice and Howard 1951) and is
generally accepted as the major agent of gene flow
among populations (Wiklund 1996). Dispersal by
reproductive adults, if it occurs, has consequences for
the lifetime reproductive success of individuals
(Clutton-Brock 1988, Newton 1989). Because of its
importance for interpopulation genetic structure and
local population dynamics, we focused our analyses on
patterns of natal dispersal, rather than breeding
dispersal.
We do not address in detail the variety of reasons,
proximate and ultimate, why an animal might disperse
from its natal area, the factors that directly determine
mortality during dispersal, or year-to-year variation in
dispersal success. However, given that animals do
disperse and that the chances of dispersers successfully
colonizing a site remain approximately constant from
year to year, we do consider the nature of relationships
among distance dispersed, body mass, and other life-
history attributes of species.
METHODS
Data collection
We searched papers published between 1930 and mid-
1998 for reports of natal dispersal movements by birds
and mammals, excluding marine species and bats. Our
geographically unrestricted survey did not include
papers written in languages other than English or
unpublished theses (except in cases where raw data for
a species were made available to us). Most
documented long-distance movements are based either
on incidental observations of dispersals made by
individual animals or on much more detailed
investigations that yield frequency distributions of
dispersal distances for one or more populations. More
observations of a single long distance moved by a
dispersing animal were reported in studies of
mammals than in studies of birds.
Wide variation in study designs, objectives, and
methods of quantifying dispersal required that we
develop screening criteria for selecting dispersal data
suitable for comparative analyses. Therefore, we
accepted dispersal data only if authors:

1. reported individual dispersal distances (or
ranges) rather than means or medians only;
2. reported net (minimum straight-line distance)
rather than gross distance moved;
3. did not include observations of likely migrants
as dispersers. We accepted subsets of data if
authors provided some evidence of the
nonmigratory status of some individuals.
Distinguishing dispersal from migration is
important for several reasons. In birds,
postfledging exploratory movements may have
a function in locating future breeding sites,
locating sites suitable for overwintering, or
establishing a navigational target, all of which
could confound interpretation of dispersal
movements (Baker 1993);
4. gave single observations of long-distance
dispersal, provided the authors had evidence
that the disperser was marked at or near its
natal site and that the reported movement met
our criteria for natal dispersal movements (as
defined above); or
5. either observed breeding (or, for banding
studies, specified breeding season returns) or,
for species with delayed maturity, provided
evidence of little movement after settlement.
Many studies reporting data on natal dispersal do not
control biases in measurement, such as the decreased
likelihood of detecting a dispersing animal at the edge
of a study area. Therefore, we further screened studies
presenting distance-density distributions of dispersal,
selecting only those that did not truncate distances by
using a small study area. We did include studies
involving several small study areas with a potential for
observing movement between sites. We accepted
statements of the adequacy of site size when surveys
of surrounding areas did not extend the distribution of
distances. Some authors also included correction
factors for potential biases in their data; we used these
if provided.
For each study selected, we recorded all given
dispersal data. If raw distance-density data were
provided in tables, we recorded them, maintaining data
separately for each sex if possible. Numerical values
were also estimated from figures. Small figures with
log scales probably resulted in some measurement
error for long distances. We converted distances given
as numbers of territories or home ranges crossed to
kilometers, assuming packed circular territories of the
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mean size measured in the study. Where available, we
also recorded median, mean, and maximum dispersal
distances. For each study, we recorded how
observations were obtained (e.g., radiotelemetry,
tagging, hunter returns, etc.). Within studies, telemetry
data were chosen over single observations when both
were given. We ignored data on nondispersal
movements.
We determined mean adult body mass for each species
and sex from the original studies (if given) or from
standard references: Dunning (1993) for birds and
Silva and Downing (1995a) for mammals. We also
identified the following categorical variables as
potential ecological and life-history correlates of
dispersal distance:
1. Diet type. We classified all bird and mammal
species into three diet groups based on the
main foods consumed throughout the year:
herbivores (including seed eaters), carnivores
(including insectivores), and omnivores.
Definitions follow Schoener (1968): herbivores
ingest < 10% by volume animal matter,
whereas carnivores ingest > 90% by volume
animal matter on an annual basis. All species
ingesting 10–90% animal matter are
omnivores. We used supplemental dietary
information from Ehrlich et al. (1988) for birds
or from our source studies.
2. Social system during the breeding season. We
classified this as territorial, nonexclusive (broad
overlap between home ranges), or gregarious
(including herding and colonial species).
3. Migratory status (birds only). We classified
bird species as migrants (including partial
migrants) or residents. All of our dispersal data
for mammals came from nonmigratory periods,
so we did not include this variable for
mammals.
4. Phylogeny. To assess the influence of
relatedness on dispersal scale, we included
taxonomic family as a categorical variable in
our analyses.
The data and references used in our analyses are
summarized in Tables A1 and A2 in Appendix 1.
Data analysis
Our general approach was to use robust methods for
estimation wherever possible and to develop simple,
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empirical relationships for prediction, rather than
constructing process-based models. Dispersal is a
complex phenomenon that makes it difficult to
develop general models based on interspecific
comparisons (Paradis et al. 1998). Besides the
previously mentioned potential for sampling biases,
sample sizes differ between studies, fates of dispersers
are not always known, and dispersers of different
species may be responding to different proximate
factors. We therefore treated our data as follows.
To examine underlying similarities in dispersal
patterns between sexes and among species, we needed
to rescale the frequency distributions of dispersal
distances obtained from each study to a common unit
of measurement. We rescaled each distribution to a
ratio of the distance moved by 50% of the observed
dispersers (hereafter referred to as the median
distance). If the distribution was given in intervals, we
used the midpoint of the interval containing the
median. Subsequent analyses involving frequency
distributions used these rescaled values.
To evaluate the relationships between dispersal
distances and life-history characteristics, we used the
following methods based on median and maximum
distances moved by dispersers of each species. First,
we logarithmically transformed all body masses and
dispersal data (median and maximum distances) to
stabilize variances, linearize responses, and normalize
residuals. We then examined normal quantile-quantile
plots of all transformed data to test for departures from
normality, finding that most dispersal distance data
remained significantly non-normal after log10
transformation.
Second, we used ordinary least squares (Type I)
regression to develop all predictive allometric
relationships, because we did not have error
distributions of body mass data for most species of the
type proposed by LaBarbera (1989) in his discussion
of appropriate regression methods in allometric
analyses. Before finalizing the regressions, we
examined the influence of each observation on the
estimated regression coefficients using Cook's (1977)
distance and eliminated observations with a Cook's
distance > 1.0. All estimated coefficients derived from
allometric equations based on log10 were corrected for
bias using the methods of Sprugel (1983). Because of
departures from normality, we estimated standard
errors and confidence limits of the allometric equation
coefficients using 1000 replicate bootstrap samples of
the data. We adjusted for bias following Efron and
Tibshirani (1993). All analyses were done with S-Plus
(MathSoft 1998).

Table 1. Summary shape statistics (means ± SD) for the original frequency distributions of detected natal dispersal distances
analyzed in this study. Distributions presented here were not transformed or rescaled prior to analysis. The skewness and
kurtosis of each distribution were calculated using Pearson moment statistics. Tests of the deviation of the population of
shape parameters from a normal distribution were done using t0.05 (Sokal and Rolf 1995).

Percent with Skewness Kurtosis

n
1 mode
λ3 P λ4 P

Birds

Females 24 50.0 1.06 (± 1.37) < 0.001 2.21 (±4.88) < 0.050

Males 23 58.8 1.40 (± 0.63) < 0.001 1.85 (±2.49) < 0.002

Sexes not separated 21 40.0 1.50 (± 1.05) < 0.001 2.44 (±5.71) < 0.20

Mammals

Females 22 62.5 1.51 (± 1.08) < 0.001 2.91 (±4.09) > 0.400

Males 20 63.6 1.22 (± 1.09) < 0.001 2.09 (±3.09) < 0.001

Sexes not separated 5 50.0 0.75 (± 0.84) > 0.15 -0.21 (±2.54) > 0.50
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RESULTS

Usable dispersal data were found for 77 bird and 68 Fig. 1. Empirical distributions of distances moved by
mammal species (see Tables A1 and A2 in Appendix juveniles dispersing from their natal areas. Each data point
1). Dispersal data were reported separately by sex in represents the number of juveniles dispersing that particular
the majority of cases: 52% for birds and 76% for distance according to the original study. All dispersal
mammals. Within each class, the range of natal distances shown are scaled to the median dispersal distance
(D) for the population estimated from the data presented in
dispersal movements varied by at least two orders of
each original study. Sexes are pooled for each class. Also
magnitude for both median distances. In the case of shown for each class is the LOESS smoothed line through
birds, median distances ranged from 0.03 km for the the data (Cleveland et al. 1992) using a quadratic smoothing
House Martin (Delichon urbica) to 10 km for the algorithm.
Glaucous-winged Gull (Larus glaucescens). For
mammals, median distances ranged from 0.03 km for
the common vole (Microtus agrestis) to 129.7 km for
the gray wolf (Canis lupus). Maximum distances for
birds ranged from 1.3 km for the European Magpie
(Pica pica) to 1305 km for the Great Horned Owl
(Bubo virginianus), and, for mammals, from 0.14 km
for the prairie vole (Microtus ochrogaster) to 930.1
km for the lynx (Lynx lynx).

Distributions of dispersal distances

The distributions of the 107 dispersal distance data

sets exhibited left skewness and platykurtosis (Table
1), indicating that many dispersers cluster near the
natal areas, whereas others disperse relatively far from
their natal areas (Turchin 1998). This pattern was
significant for all class and sex comparisons, except
for mammals in studies where the sexes were not
separated. Shape statistics based on the original
distance-density distributions are highly variable,
particularly for distances dispersed by females (Table
1). This variability, coupled with different methods of
categorizing distances among studies, suggested that
further analysis of sources of variation in the patterns
of dispersal in birds and mammals was warranted.

Once distances were rescaled to units of the median

dispersal distance for each distribution, comparative
analyses revealed that the distributions were
statistically indistinguishable between sexes and
between noncarnivores (herbivores and omnivores
combined) and carnivores. In two sample
Kolmogorov-Smirnov tests, the sex-based values were
D = 0.042, Pα=0.05, 100 > 0.5 for birds and D = 0.023,
Pα=0.05, 100 > 0.5 for mammals. The noncarnivore vs. Two broad patterns were evident in these pooled data.
carnivore values were D = 0.015, Pα=0.05, 100 > 0.5 for First, rescaled dispersal distance distributions for bird
birds and D = 0.029, Pα=0.05, 100 > 0.5 for mammals. and mammal species were very similar in shape (Fig.
Consequently, for both classes we pooled distributions 1). Second, for both classes, the rescaled dispersal
for different sexes and diet types. distributions showed a strong decline in frequency
with increasing distance from the natal area (Fig. 1). In
both birds and mammals, most detected dispersal

distances were less than 3 median distances from the
natal area; only 9 of 35 bird species and 2 of 19
mammal species showed juveniles dispersing farther
than 10 median distances. At least part of the decline
in the numbers of birds and mammals was probably
due to the reduced likelihood of detecting animals far
from the natal area. Although we attempted to screen
out data where this problem was clearly evident, subtle
sampling biases of this type were more difficult to
identify. We therefore treated the rescaled distributions
as estimates of minimum dispersal distances only, and
not as estimates of "true" but unknown dispersal
distances.
Rare long-distance dispersals often appear at the
extreme right of frequency distributions of dispersal,
although their effects on determining the dispersal
scale of species are usually difficult to take into
account (Turchin 1998). To test for the presence of
"fat tails" in the rescaled distributions, we used
polynomial regression to examine the significance of
the coefficients (b0, b1, b2) in the relationship
ln(dispersers)x = b0 + b1x + b2x2, where x is distance
dispersed. We found that, in both birds and mammals,
the b2 coefficients were significantly greater than 0: for
birds, this value was t0.05, 19 = 8.93, P < 0.0001; for
mammals, t0.05, 12 = 7.67, P < 0.0001. This confirmed
that the tails were fatter than expected under an
assumption of a Gaussian distribution (Turchin 1998).
Although more detailed statistical analysis of the tails
is not warranted here because of potential biases in the
original data, this result is consistent with a dispersal
pattern created by highly variable dispersal distances
that also includes rare long-distance dispersal events.
Allometric scaling of dispersal distances
The frequency of natal dispersal (i.e., the proportion of
individuals dispersing) is female-biased in birds and
male-biased in mammals (Greenwood 1980), leading
to the expectation that female birds and male
mammals may also disperse farther than their
counterparts of the opposite sex. We could not find
strong enough evidence for these hypotheses in our
data to warrant separate analyses for males and
females. There are high probabilities that median and
maximum dispersal distances show no significant sex
bias. Sign tests comparing distances dispersed by
females and males of the same species showed that,
for birds, median distances were P = 0.402, n = 16 and
maximum distances were P = 0.254, n = 9, whereas,
for mammals, median distances were P = 0.613, n =
12 and maximum distances were P = 0.06, n = 8.
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However, it is possible that the lack of significance in
our findings for the maximum distance comparison in
birds and mammals was due to small sample sizes.
Except where indicated below, we pooled the sexes
together for the remainder of our interspecific
comparisons.
Do dispersal distances scale with body mass and other
ecological factors that may determine the density and
dispersion of breeding sites across the landscape?
Based on our initial expectations, we tested the
importance of average adult body mass M in
explaining variations in median dispersal distances
(Dmedian) and maximum dispersal distances (Dmax) in
birds and mammals. Both Dmedian and Dmax increased
significantly with increasing body mass in both
classes, and the variation in dispersal distance
explained by body mass was higher for maximum
dispersal distance in mammals (Table 2). By
comparing bootstrapped confidence intervals for each
measure (median vs. maximum distance), we found
that the estimated slopes of these relationships did not
differ at α = 0.05 between either measure within
classes or each measure between classes. Nevertheless,
the intercepts between measures differed significantly
for each class. The estimated maximum distance that
dispersing individuals of a species with a given body
mass were expected to move was 4.9 and 5.8 times the
median dispersal distance in birds and mammals,
respectively (based on ratios of intercepts). Over the
range of comparable body masses, ratios of predicted
dispersal distances for birds and mammals combined
vary between 5.1 and 9.5 (median distance) and 5.0
and 56.0 (maximum distance). Overall, birds dispersed
between 0.5 and 1.5 orders of magnitude farther than
mammals of equivalent body mass. Although body
mass was a significant predictor of both median and
maximum dispersal distances for mammals, it was
clearly weak in predicting maximum dispersal distance
for birds.
When we separated species within classes by diet type,
estimated allometric relationships were consistently
stronger for carnivorous species than for herbivores or
omnivores (Figs. 2 and 3). For herbivorous and
omnivorous birds, the percentage of unexplained
variance in median or maximum dispersal distance
ranged between 86% and 98% after accounting for
body mass. The slopes of the allometric regressions
were not significantly different from 0 for either diet
type or dispersal measure in herbivorous and
omnivorous birds (P > 0.154 for all slopes), whereas
the slopes were significantly different from 0 for
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carnivorous birds (P < 0.0005 for both median and 1.21; maximum, 0.32–0.77), although there was
maximum distances). Furthermore, slopes for considerable residual uncertainty about the slope (Fig.
carnivorous birds did not differ significantly from 0.75 2).
(bootstrapped 95% confidence range for median, 0.49–

Table 2. Summary of equations converting body mass M (in kg) to type of dispersal distance (in km) in birds and mammals.
Shown are separate Type I regression coefficients (± SD) and regression statistics for species of each diet type ("C" =
carnivores, "H+O" = herbivores and omnivores pooled, and "Combined" = all diet types combined).

Class Dispersal distance type Trophic type Equation df r2(adjusted) P

Birds Median C 36.4 (± 1.33) M0.62 (± 0.27) 12 0.24 0.0440

2.1 (± 1.76)
H+O 24 0.04 0.3300
M0.18 (± 0.18)

13.1 (± 1.47)
Combined 36 0.32 0.0001
M0.63 (± 0.15)

Maximum C 199.5 (± 1.38) M0.59 (± 0.13) 34 0.33 0.0003

H+O 36.4 (± 1.55) M0.14(± 0.15) 41 0.02 0.6600

Combined 73.3 (± 1.31) M0.34 (± 0.10) 75 0.09 0.0030

Mammals Median <

C 3.45 (± 1.07) M0.89 (± 0.03) 10 0.88
0.0001

<
H+O 1.45 (± 1.05) M0.54 (± 0.01) 18 0.74
0.0001

Combined 2.04 (± 1.32) M0.67 (± 0.09) 28 0.42 0.0010

Maximum <
C 40.7 (± 1.41) M0.81 (± 0.14) 48 0.73
0.0001

H+O 3.31 (± 1.17) M0.65 (± 0.05) 15 0.75 0.0001

<
Combined 6.46 (± 1.23) M0.68 (± 0.08) 63 0.72
0.0001

For mammals, the unexplained variance in median and
maximum dispersal distances after accounting for
allometric relationships was less than 32% for all
regressions. Analysis of covariance indicated that the
intercepts of the relationships for carnivorous
mammals differed significantly from those of
herbivores and omnivores (P < 0.03), although the
slopes of the regressions did not differ. We therefore
pooled our data for herbivores and omnivores (Fig. 3).
Dispersal distances of mammalian carnivores were
considerably longer than those of herbivores and
omnivores. The predicted ratios of carnivore dispersal
distances to herbivore/omnivore distances ranged
between 1.2 and 4.5 (median distance) and 7.1 and
18.3 (maximum distance) for mammals of equivalent
body mass. As in birds, the slopes of the regressions
for carnivorous mammals did not differ significantly
from 0.75 (bootstrapped 95% confidence range for
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median, 0.65–1.11; maximum, 0.64–1.10), although 0.75 (bootstrapped 95% confidence range for median,
the slopes for both measures calculated for the 0.39–0.70; maximum, 0.56–0.72).
combined herbivore and omnivore data were both <

Fig. 2. Allometric relationships in birds between body mass (M), median dispersal distance (Dmedian) on the left, and
maximum dispersal distance(Dmax) on the right. Solid triangles represent data points for herbivores, open circles represent
data points for omnivores, and solid circles represent data points for carnivores. Significant Type I regressions (—) are shown
for carnivores only (see Table 2 for equations), along with their 95% confidence bands (····) and 95% prediction bands (----).
Sexes or other categorical variables are not differentiated in this figure.
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Fig. 3. Allometric relationships in mammals between body mass, median dispersal distance on the left, and maximum
dispersal distance on the right. Symbols are as in Fig. 2. Significant Type I regressions are given for herbivores and
omnivores (pooled) and for carnivores (see Table 2 for equations). Sexes and other categorical variables are not
differentiated. Note the difference in x-axis scale between this figure and Fig. 2.

We examined the relationships between our two
measures of dispersal distance (median and maximum)
and the categorical life-history variables after we
removed the influences of body mass and diet type
(Table 3). Except for family and migratory type in
birds, we found little evidence for the effects of these
variables on either median or maximum dispersal
distances in birds or mammals (Table 3). We found
that observed maximum distances dispersed by species
of Corvidae, particularly Gray Jay (Perisoreus
canadensis) and Magpie (Pica pica), and Fringillidae,
namely Song Sparrow (Melospiza melodia), Savannah
Sparrow (Passerculus sandwichensis), and White-
crowned Sparrow (Zonotrichia leucophrys), were
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shorter, averaging 72% less than the predicted value.
In contrast, those of the Laridae, including the lesser
Black-backed Gull (Larus fuscus), the Glaucous-
winged Gull (L. glaucescens), the Ring-billed Gull (L.
delawarensis), and the Herring Gull (L. argentatus),
and the Motacillidae, particularly the White Wagtail
(Motacilla alba) and the Yellow Wagtail (M. flava)),
were longer than expected, averaging 60.7% farther
than the predicted value based on the general body
mass-diet type relationship for birds. In addition, the
maximum distances dispersed by migratory bird
species were significantly farther than for resident
species (Table 3); class averages for migratory species
were 28.4 km, n = 39, and, for resident species, 24.3
km, n = 34. Our data provided no clear evidence that
social system was a correlate of the distance of
dispersal in either birds or mammals, although, in
general, low power precluded eliminating this variable
from further study (Table 3).

Table 3. Effects of categorical variables on median and maximum dispersal distances of birds and mammals after removing
the effects of body mass and diet type. Shown are all main effects and significant interactions. Effects significant at α = 0.05
are marked with an asterisk (*). Comparisons with too few observations for analysis are indicated by an ellipsis ("...").

Median dispersal Maximum dispersal

Class Variable
F d.f. P Power F d.f. P Power

Birds Family 2.21 9,27 0.05* 0.88 0.64 11,77 0.77 0.35

Migration 0.01 1,27 0.93 0.10 0.68 1,77 0.41 0.15

Social
0.08 2,27 0.27 0.12 0.10 2,77 0.91 0.11
system

Migration x
social ... ... ... ... 191.20 22,77 0.046* 0.97
system

Mammals Family 0.22 10,17 0.99 0.98 1.76 15,48 0.20 0.64

Social
2.14 1,17 0.16 0.45 0.11 2,48 0.89 0.12
system

Family x
0.02 1,17 0.89 0.10 0.80 30,48 0.73 0.41
social system

By studying patterns of residuals after the influences 0.49 for mammals.

of body size and diet type had been removed, we
tested for two possible trends in our data. First, we
searched for systematic biases introduced by the
method of obtaining dispersal data. We found no
evidence that the observation method biased the results
in either birds or mammals. One-way ANOVA yielded
a median distance of F0.05, 3,347 = 1.17, P = 0.33 and a
maximum distance of F0.05, 3,74 = 1.42, P = 0.51 for
birds, and a median distance of F0.05, 2,10 = 1.72, P =
0.20 and a maximum distance of F0.05, 2,64 = 0.72, P =
Second, because birds are more mobile than mammals,
we considered the possibility that their observed
dispersal distances might also be more variable than
those of mammals. When we looked at how the
observed dispersal distance values were dispersed with
regard to their predicted values between classes and
diet types, we found no evidence that this was the
case. In fact, it is possible that dispersal distances for
mammals may be relatively more variable than those

for birds: paired t tests of coefficients of variation
yielded values of t0.05(2),3 = 1.95, P = 0.15.
Fig. 4. Relationship between the median dispersal distance
(Dmedian) and the maximum dispersal distance (Dmax)
observed in birds and mammals. Birds are represented by
open circles and mammals by solid circles. Also shown is
the fitted Type I regression line (——) for the pooled data,
the 95% confidence band (····), and the 95% prediction band
(----) for the relationship. The sexes are not differentiated.
Because we could estimate the median and maximum
dispersal distances for a given species from the body
mass and diet type relationships in Figs. 2 and 3 and
Table 2, we wondered if we could also estimate the
median distance dispersed from the maximum
distance, and vice versa. To examine this, we chose
only those species of birds and mammals for which we
had both a maximum dispersal distance observation
and a median dispersal distance. As expected, both
classes showed a significant relationship between
maximum and median distances, and, as seen in Fig. 4,
Dmax = 9.77 (± 1.23 SD) x Dmedian = 0.98 (± 0.08 SD),
df = 59, adjusted r2 = 0.73, P = 0.0001. For both birds
and mammals, the slope of the line was not
significantly different from a 1:1 relationship, although
birds had a higher intercept than mammals.
An empirical probability model for the
minimum distance of dispersal
To infer the minimum probability that a dispersing
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animal could disperse a given distance using data on
its body mass, diet type, and the relationships in Table
2 and Figs. 2, 3, and 4, we derived a simple
phenomenological model based on the empirical
density-distance curves in Fig. 1. Our intention was to
develop general relationships for estimating the
minimum probability that a given animal would
disperse beyond a particular distance. For the range of
species that we examined, potentially there were many
varied mechanisms that could govern the decision of a
dispersing individual to settle at a site. We chose a
conservative dispersal model based on negative
exponential distribution as the simplest and most
general description of the data. This distribution
assumed that, over a large area and a sufficiently long
time, the probability of predicting the dispersal
distances of individuals could be approximated by a
Poisson process. Other process-based models derived
from alternative hypotheses to determine where
dispersers settle have also been developed (Taylor
1980, Miller and Carroll 1989, Caley 1991, Turchin
1998). In many cases, these models do not describe
dispersal patterns any better than does the negative
exponential, even for single species (Miller and
Carroll 1989, Caley 1991). If X is the potential
distance dispersed by an individual originating in a
natal area, with an expected distance of θ, then the
distribution function of X
(1)
with the useful property that the probability that X
exceeds some value x is given by
(2)
when x > 0 (Hogg and Tanis 1997).
For each class, we fit separate negative exponential
functions to two distributions. The first, called the
"base" estimate because the source data probably
underestimated the true pattern of dispersal due to
unknown distance-weighted sampling biases, was the
mean value of the empirical distribution given at each
distance unit in Fig. 1. This value was Pr(distance
dispersed > D) = e-(D/1.64), r2 = 0.99 for birds, and
Pr(distance dispersed > D) = e-(D/1.5), r2 = 0.99 for
mammals. The second was a "corrected" estimate of

mean dispersal made using mean correction factors for
birds and mammals based on the factors estimated for
individual species by Porter and Dooley (1993). In this
case, the values were, for birds, Pr(distance dispersed
> D) = e-(D/2.6), r2 = 0.94, and, for mammals,
Pr(distance dispersed > D) = e-(D/2.06), r2 = 0.96. The
resulting pairs of probability curves are shown in Fig. 5.
Fig. 5. Estimated probabilities that the minimum distance
moved by dispersing birds or mammals exceeds a particular
number of median distance units (D) based on negative-
exponential models fit to the empirical distance density data
(see An Empirical Probability Model for the Minimum
Distance of Dispersal). Fitted parameters are presented for
"base" assumptions and for estimated distributions using
distance-weighted sampling correction factors representing
"corrected" assumptions. The horizontal dashed lines
indicate 0.05 probabilities that dispersing juveniles will
move farther than the distance indicated by the matching
vertical dashed lines.
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The primarily herbivorous red squirrel (Tamiasciurus
hudsonicus) and the carnivorous marten (Martes
americana) are good examples of how these
probability models can be used in combination with
allometric equations to yield estimates of the minimum
probability that dispersers of selected mammal species
will disperse a given distance. Female red squirrels
have an average body mass of 0.199 kg (Appendix 1,
Table A2), and the equations in Table 2 predicted
median and maximum dispersal distances of 0.82 and
11.0 km, respectively. Using the "base" probability
model equation for mammals given above, we found
that the estimated probability of dispersing farther than
11.33 median distance units was less than one in a
thousand. By multiplying the estimate of the median
distance dispersed for this species (0.82 km) by this
estimate of the number of median distance units
(11.33), we found, using our "base" assumptions, that
dispersing females of this species had a small
probability of settling farther than 9.29 km. Similar
calculations for marten, which has a body mass of 1.04
kg, yielded an estimate of 40.5 km as the threshold
distance beyond which dispersers have only a small (P
< 0.001) probability of successfully dispersing. For
marten we also had one long-distance dispersal
observation of 61 km (Appendix 1, Table A2). Given
the conservative assumptions we used, our estimate of
a probable "base" threshold long-distance dispersal
range for marten was, as expected, smaller than this
observation. The equivalent threshold long-distance
dispersal range calculated using the "corrected"
probability model was 51.0 km for this species.
How well do the empirical minimum-probability
negative exponential models predict the observed
maximum extent of dispersal distances in birds and
mammals? The dispersal distance data set used to
estimate parameters for the empirical models (Fig. 1)
did not include the maximum long-distance dispersal
movement used to develop the allometric models
(Figs. 2 and 3). Hence, we compared the probabilities
of observing dispersal distances greater than those
estimated from the empirical model with the
proportion of observed maximum distances from our
data that did exceed the predicted maximum distance
(Fig. 6). As expected given the evidence of "fat tails"
in the distributions of dispersal distances, all the
models underestimated the frequency of long-distance
dispersal events, i.e., all points were above the 1:1
line. This was true for both classes even when the
estimated maximum extent was large enough that the
predicted probability of observing even longer
distances became very small. For this reason, the

models are best applied in cases where researchers and
managers wish to minimize the risk of overestimating
dispersal probabilities, for example, for endangered or
highly habitat-specific species.
Fig. 6. Comparison of the probabilities that dispersers
would disperse farther than a given distance (D), estimated
using the negative-exponential models with the percentage
of observed maximum dispersal distances in birds and
mammals from our data that exceeded the predicted
distance. Error bars represent 95% confidence intervals
about the mean values, calculated using 1000 replicate
bootstrap samples of the data. Separate comparisons are
shown for the "base" probability models, which are
represented by open symbols, and for the "corrected"
probability models, which are represented by solid symbols.
Also shown are the 1:1 lines indicating an identity between
model predictions and observations.
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DISCUSSION
We found that a significant proportion of the variation
in the distances dispersed by juvenile birds and
mammals could be explained by differences in body
mass and diet type, despite known differences among
species in terms of reproductive ecology and movement
capability. Within a class, large species dispersed
farther than small species. In comparisons of bird and
mammal species of similar body mass, the dispersal
distances of carnivorous species were significantly
longer than those of noncarnivorous species. In
addition, the dispersal distances of carnivorous species
of both birds and mammals grew at an increasing rate
with increasing body mass. Our finding that the shapes
of distributions of dispersal distances were similar
among species and classes suggested that the
probability of individuals dispersing different distances
could be estimated even for poorly known species. For
many research and management situations, it would be
helpful to calculate the approximate likelihood that
organisms could disperse particular distances between
required habitats. One way to do this is to combine a
predictive allometric relationship for estimating long
distances of dispersal with a general phenomenological
model for predicting the minimum probability of
dispersing at least that far. We think that a primary
benefit of our models lies in their ability to identify
attainable distances for populations and species using
easily acquired characteristics of species (attributes of
diet and body mass) if detailed dispersal data are
lacking. Our models can be used to identify dispersal-
limited species and thus focus attention on the
characteristics of the landscapes needed to maintain
viable populations of these species.
Consequences of body size and diet type for
dispersal scale
Do patterns of natal dispersal reflect an underlying
relationship among macroecological variables such as
habitat productivity, patterns of resource use, and the
processes that determine how far animals disperse?
Among bird and mammal species, differences in the
coefficients of the allometric regressions for different
diet types were found to be consistent with expected
differences in the density and spacing of resources.
Intercepts for the equations for carnivores predicted
dispersal distances over two orders of magnitude
farther than those of herbivores and omnivores of
equivalent body mass. The slopes of the relationships
for carnivores were consistent with the ¾ scaling
expected for size-dependent resource utilization

relationships (McNab 1963, Peters 1983, West et al.
1997). Slopes for herbivores and omnivores were
smaller than those for carnivores. We remain cautious,
however, when relating causal processes to these
statistical differences between allometric slopes,
treating them instead as summary descriptions of the
minimum distances of dispersal among species. We
think that the complex spatial and temporal
interactions that determine when animals move, their
risks of mortality while dispersing, and the processes
driving habitat availability must all help explain
differences in dispersal patterns based on resource
utilization (see also Speculation section below).
Methods of fitting allometric equations differ in their
assumptions and interpretation of results. Reduced
major-axis (Type II) regression provides a less biased
estimate of the underlying functional relation as
described by an allometric equation (LaBarbera 1989),
and some authors base their allometric analyses on
Type II regression (Silva 1998). Error distributions of
body mass data for species were provided only
occasionally in the sources we used. This prevented us
from using Type II regression in most analyses.
However, our primary goal was to use the equations
for predicting expected dispersal distances, given body
mass and other criteria. For this purpose, Type I
regression was appropriate (LaBarbera 1989).
The spatial scale of dispersal is a primary determinant
of its functional role in the dynamics of populations,
even in sedentary species of birds (Koenig et al. 1996,
Martin 1998). The frequency distribution of dispersal
distances is key to calculating the probability that
individuals can move through a given landscape
(Merriam 1998). Our models can be used to estimate
minimum threshold values of the likelihood that
dispersing individuals can move particular distances.
Despite the assumptions in our models, we think that
they can be used to generate distributions of dispersal
distances for species whose dispersal parameters are
poorly known. Our models have heuristic value in
estimating minimum distances between suitable
habitats for selected species, e.g., conservatively,
female red squirrels (T. hudsonicus) have a strong
probability (P < 0.001) of successfully dispersing at
least 9.2 km, whereas marten (M. americana) have the
same probability of successfully dispersing at least
40.5 km; details of these calculations are provided in
the Results section. However, our models cannot be
used to calculate directly the rates at which species
could expand their ranges. Rare, long-distance
dispersal events influence the overall pattern and scale
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of dispersal in both birds and mammals. For example,
the leptokurtic, fat-tailed shapes of the distributions in
Table 1 suggest highly variable dispersal distances
among individuals of a given species, with a few
individuals dispersing long distances relative to the
median of the population. Such rare long-distance
events are difficult to observe but are important for the
spread of species (Kot et al. 1996), as well as for
preventing inbreeding in small populations (Mills and
Allendorf 1996). Simple probability density models
such as the negative exponential, while capturing the
average behavior of the population of dispersers,
significantly underestimate the potential for range
expansion in many organisms (Kot et al. 1996).
By examining the predictive nature of intra- and
interspecific patterns of dispersal distances in both
birds and mammals, we can extend results from other
research. Earlier work by Peters (1983) found positive
relationships among body mass, speed of locomotion,
and maximum distance of migration for both birds and
mammals. The general allometric patterns that we
obtained for measures of natal dispersal in mammals
are similar to those obtained by Van Vuren (1998) for
median dispersal distances of 40 species of North
America mammals and by J. O. Wolff (unpublished
data) for long-distance movements in 74 species of
mammals. We obtained slightly shorter median
dispersal distances for herbivorous mammals than did
Van Vuren (1998), although the differences are within
95% confidence intervals as calculated from his
results. In their analysis of natal dispersal patterns in
75 terrestrial bird species in the UK, Paradis et al.
(1998) found that body mass was a significant
covariate of mean and median dispersal distances, but
they did not attempt to develop predictive
relationships. Body mass, diet type, and other factors,
in part, determine dispersal distance. Thus, these
factors assist in linking broad population-level
consequences of dispersal (e.g., lifetime fitness,
probabilities of habitat occupancy, genetic relatedness)
with life-history attributes of species and their needs
for resources, including the distribution and
availability of these resources within and among
habitats (Brown and Maurer 1989, Holling 1992,
Martin 1998). Specific predictions about the dispersal-
mediated consequences to population dynamics that
are created by future perturbations of habitats are less
easy to develop using our results. Factors such as
density-dependent effects on dispersal rates, mortality
risks, and the availability of unoccupied areas in
habitats result from interactions among behavioral
processes below our level of analysis.

Life-history correlates of dispersal scale
We were surprised to find little evidence that the
median or maximum distances of natal dispersal in
birds or mammals were related to broad categories of
life-history strategies. Only in birds were taxonomic
relatedness and migratory status found to be related to
maximum distance dispersed. In general, our findings
paralleled those of Paradis et al. (1998) despite the
differences in our statistical approach. Of particular
interest are the differences in dispersal distances
between migratory and resident bird species, with the
former dispersing farther. Resident species probably
incur different costs and benefits from dispersing than
do long-distance migrant species (e.g., neotropical or
transequatorial migrants) and may be less
opportunistic in their choice of a site at which to settle
(Paradis et al. 1998). However, our questionable
ability to separate postfledging exploratory movements
(including dispersal) from migratory movements in
our data renders this interpretation provisional.
Furthermore, our tests are generally not very effective
at distinguishing putative differences in dispersal
patterns attributable to other life-history factors (range
of β: 0.05-0.48). We therefore suggest that allometric
scaling of dispersal among species as presented in our
study should be viewed primarily as representing
synthetic relationships that integrate many, sometimes
conflicting, fine-scale behavioral and ecological
processes rather than indicating their lack of relevance.
Several syntheses showed that dispersal rates and
distances were often male-biased in mammals and
female-biased in birds (Greenwood 1980, Wolff 1993,
Clarke et al. 1997). In our analysis, evidence of a
distinction in median or maximum dispersal distances
between sexes of the same species was too weak to
make it worthwhile to treat the genders separately.
Possible explanations for our failure to detect expected
differences between genders in median and maximum
distances include small sample sizes (only 16 species
of birds and eight of species of mammals met our data
criteria), high variance (in part, because observations
of maximum distances dispersed by individual animals
are often serendipitous), and the fact that our dispersal
data did not come from a random sample of species.
Bird species dispersed much farther than mammal
species of equivalent body mass for both median and
maximum dispersal distances. This result is intuitive,
because birds are more vagile than nonvolant
mammals.
It was even more surprising that we did not find
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evidence that the range of variation in median or
maximum dispersal distances was greater for birds
than for mammals, after accounting for body mass and
diet type. It is possible that differences in researchers'
ability to detect dispersing birds and mammals may
have confounded this result. However, not only did we
fail to find a detectable effect of observation method
(e.g., radiotelemetry, band-resight, etc.) on the residual
variation about the regressions, but we also screened
both sets of data to ensure that the effects of small
study areas were reduced or eliminated. This apparent
similarity in range of variation for dispersal distances
between the two classes may reflect interclass
similarity in types of interactions between dispersing
animals and their environment that determine their
probability of settling. Our inability to separate clearly
the effects of physiological limits, mortality factors,
and variation of habitat quality on the scale of
dispersal constrains our ability to infer how those
interactions might operate.
Martin (1998) posed basic questions about patterns of
dispersal: "how, where, how far, when." Our
interspecific study identified predictive relationships
underlying the "how far" question. However, other
data are needed from species-specific studies of
dispersal. From our perspective of dispersal processes
and landscape design, an important gap in knowledge
is the lack of information about the survivorship of
dispersing animals. Simulations by Henein and
Merriam (1990) showed the potential importance of
survivorship during dispersal movements, and there
are empirical data from several studies involving the
dwarf mongoose, Helogale parvula (Waser et al.
1994), the Blue-breasted Fairy Wren, Malurus
pulcherrimus, and the White-browed Babbler,
Pomatostomus superciliosus (Brooker et al. 1999), and
the Northern Spotted Owl, Strix occidentalis (Miller et
al. 1997). However, studies of dispersal in several
grouse species did not indicate a significant survival
cost to dispersing juveniles (Martin 1998). We agree
with Merriam (1998) that studies of habitat-specific
survivorship during dispersal periods are required to
assess rates of exchange between fragments in
landscape-habitat models.
SPECULATION
In many vertebrates, body mass is closely correlated
with ecological variables that affect the allocation of
space and nutritional resources, e.g., home range area
(McNab 1963, Harestad and Bunnell 1979, Holling
1992), local population density (Silva and Downing

1995b), and geographic range (Brown and Maurer
1989). We recognize that there are still substantial
problems related to the derivation, interpretation, and
predictive power of allometric scaling when applying
it to interspecific ecological analyses.
First, whereas allometric relationships provide a
concise description of the percentage changes of one
structural character (usually body mass) with other
life-history characteristics, their functional
interpretation remains unclear. At root, this uncertainty
is founded upon the question of whether a pervasive
causal process underlies allometric patterns or whether
interspecific allometries arise as mere statistical
epiphenomena out of syntheses of intraspecific
patterns (Cates and Gittleman 1997). On the one hand,
West et al. (1997) propose that constraints on energy
allocation within and among organisms in a phylogeny
are the fundamental explanation for interspecific
allometric relationships (including macroecological
ones such as home range size and patterns of
population density). On the other, Koslowski and
Weiner (1997) argue that, because the target of
evolution is individual variation within a species,
interspecific patterns may combine different
evolutionary solutions to the problem of energy
allocation. Consequently, they think that interspecific
allometries have only weak biological significance.
Second, analytical methods for calculating allometric
exponents can be biased (LaBarbera 1989),
particularly in their assumption that the processes
being compared are similar across the taxa and body
mass ranges under consideration (Prothero 1986).
Thus, inferences about the meaning of exponents
relating ecological patterns to body mass must
carefully consider whether underlying processes are
truly comparable among the species included (Holling
1992).
Third, by the nature of their derivation, scaling
relationships (e.g., exponents) have little capability to
discriminate across the broad classes of stochastic or
conditionally stochastic processes that generated them.
Additionally, because they are implicitly conditional
(i.e., they arise from studies conducted under specific
conditions), using them to forecast patterns is
problematic if future conditions are expected to differ
substantially from those under which the original
relationships evolved.
Despite these difficulties, broad cross-taxon
relationships reveal characteristics of processes that
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are not always apparent from studies of individual
species. If the ultimate evolutionary function of
dispersal is to improve reproductive success (Sinclair
1992) and if interspecific allometric relationships
capture the responses of species to patterns of resource
availability at different scales (Holling 1992), then a
species' dispersal capability may be linked to the
spatial and temporal grain as well as the extent of the
resources required for survival and reproduction.
Generally, interpretations of an animal's areal
decisions about utilization of resources are related to
the density of resources at the scales of individual
foraging items, patches, and their seasonal integration
at the larger scale of a home range (Holling 1992,
Peterson et al. 1998). Our finding of structurally
similar relationships between body mass and the
spatial scale of dispersal in birds and mammals
suggests that an individual's dispersal decisions have a
similar ecological basis, serving to locate the animal in
habitats that are likely to provide resources needed for
its long-term reproductive success. We suggest that, at
the large spatial and long time scales of dispersal
phenomena, the summary properties of dispersal
processes that determine their spatial extent may be
derived from interactions between behavioral and
morphological variables of species and their linkages
to the dynamics of resource availability in landscapes.
The results of our study can be used to develop and
assess potential solutions for problems involving
habitat management and landscape planning. We
suggest two broad types of applications and offer
examples from forests. First, the basic parameters of
the frequency-distance distributions are fundamental to
assessing the potential effects of habitat loss and
fragmentation on selected species. Many conservation
issues involve natal dispersal because it is a primary
mechanism for colonization and genetic exchange
between populations (Hedrick 1996). As well,
population declines in many species have been linked
directly to loss and fragmentation of habitats
(Robinson et al. 1992) and indirectly to reduced
interpatch dispersal (Whitcomb et al. 1981, Fahrig and
Merriam 1985, Doak et al. 1992, Pulliam et al. 1992,
Lamberson et al. 1994, Schumaker 1996). For species
in which preserving key habitats is a management
priority, researchers and managers can combine the
allometric equations with the probability model to help
assess the ability of alternative configurations of
habitat patches to provide adequate connectedness
among populations (Van Vuren 1998).
Managers must have a knowledge of dispersal because

it is a critical variable for modeling the effects of
landscape change on the long-term viability of
metapopulations (Beissinger and Westphal 1998).
Concerns about habitat fragmentation and landscape
design are based, in part, on the ability of wildlife to
disperse between the blocks of habitat types that they
require (Schumaker 1996, Fahrig 1997). Long-term
population declines of forest-dependent species are
thought to occur because large-scale forest harvesting
changes habitats by destroying old-growth forest,
reducing the number of areas of contiguous mature
forest, and eliminating many structural components in
stands (Bunnell and Kremsater 1990, Hansen et al.
1991, Spies et al. 1994). Forestry practices can alter
and fragment habitats over spatial and temporal scales
important for population survival, e.g., 10 m–1000 km
and 1 month–1 century (Bunnell and Huggard 1999).
These scales place a priority on identifying the
equivalent extent of population movements that will
reduce the risk of population loss. Our study provides
estimates of dispersal distances and can be used to
predict the effects of fragmentation and alteration of
habitats on individual species.
The second broad type of application is at the level of
communities in managed landscapes. Our results can
be used to help identify dispersal-limited species that
may face long-term risks from large-scale habitat
alterations. The allometric relationships that we
present can be used to estimate dispersal distances for
each species of bird and mammal in a community and
assess their potential vulnerability to fragmentation
based on dispersal capability. Loss and fragmentation
of forests, especially late-seral stands, is touted as a
critical issue in the management of temperate forests
(Lehmkuhl and Ruggiero 1991). Predicting the effects
of habitat changes due to forest practices on
communities is contentious because of the biological
interdependence of forest ecosystems; forestry and
land-use practices create several changes
simultaneously (Bunnell 1999). To meet the habitat
needs of the community of species dwelling in late-
seral stands, managers must consider the amount of
late-seral forest that is retained, the size of forest
patches, and the distances between forest patches. It is
difficult to optimize the spatial distribution of later
seral forests for the suite of species that depends on
them because of the complex relationship between
habitat area, patch size, and distance between patches
(Harris 1984). Our models can be used as a planning
filter for assigning priorities to conservation initiatives
and more directly evaluating management options for
these vulnerable species. By examining dispersal
Conservation Ecology 4(1): 16.
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capability among the species of a community,
members of the community can be identified that are
potentially more vulnerable to habitat fragmentation
than others. Such analyses can be used to plan
landscapes of managed forests over the long term,
supporting frequently used theoretical arguments for
the inclusion of broad-scale movement patterns in the
management of wildlife habitat (Schumaker 1996,
Beissinger and Westphal 1998). By combining
information on patch size with data on distances
between patches, forest managers can determine which
forest-dwelling species are vulnerable in existing and
future landscapes.
Modeling of dispersal distances is useful at the scale of
landscapes in assessing the connectedness between
habitats for the species and populations they host.
Such analyses must include detailed species-specific
habitat relationships (Brooker et al. 1999). Besides
fragmentation, managers should consider processes
that occur within the matrix between habitat
fragments. The number of individuals dispersing a
particular distance depends on the survivorship of
dispersing animals as they travel through this matrix.
However, there is little information on the landscape
features that increase the risks of mortality for
dispersing animals (Beissinger and Westphal 1998).
To facilitate dispersal and reduce risks to vulnerable
species, it is essential to maintain the quality of the
habitat encountered by dispersing animals. Two key
questions are central to issues of forest fragmentation:
how hostile is the intervening habitat to a species and
for how long is it hostile? In forest landscapes
modified by logging, managers could decrease the
"hostility" of the matrix by maintaining structural
heterogeneity within cutblocks (Franklin et al. 1997).
By providing suitable cover in cutblocks, survivorship
of dispersing individuals could increase, which would
enhance dispersal between fragments. Appropriate
silvicultural practices in the matrix between forest
fragments could reduce the vulnerability of species to
habitat fragmentation.
Responses to this article can be read online at:
http://www.consecol.org/vol4/iss1/art16/responses/index.html.
Acknowledgments:
We thank Fred Bunnell, Dave Daust, Glen Dunsworth, Sue
Glenn, David Huggard, Kathy Martin, Doug Runde, and Ian
Thompson for helpful discussions during the development of
this paper. Fred Bunnell, Clay Elder, Neil Surrey, and Elke
 Conservation Ecology 4(1): 16.
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Wind assisted us with the literature search. Comments by
the Forest Ecology Group at SFU, Sue Glenn, David
Huggard, C. S. Holling, Phil Taylor, and three anonymous
reviewers greatly improved earlier versions of the
manuscript. Research funding was provided by a Forestry
Canada Green Plan research grant to ASH, KL, and Fred
Bunnell, as well as Forest Renewal British Columbia, and
an EcoResearch Doctoral Fellowship to GDS. This is
research contribution R-33 from the Centre for Applied
Conservation Biology.

APPENDIX 1
Sources of dispersal data for birds (Table A1) and mammals (Table A2) used for analysis. Species are listed in
taxonomic order. For each study, the type of data extracted is coded as "S" (single observation only—generally a
maximum natal dispersal distance) or "D" (distance-density distribution). Categorical life-history variables shown
here are: (1) diet type: H, herbivore; O, omnivore; C, carnivore; (2) social system in breeding season: T,
territorial; N, non-exclusive territories or neighborhood; G, gregarious; and (3) migratory status of the population
or species: M, migratory; R, non-migratory. If sample sizes (n) are given, they refer to the number of dispersing
animals meeting our criteria that were reported in the study; otherwise, a " ... " is given. Where possible, data for
males (m) and females (f) are shown separately. Data from studies in which the sexes were not differentiated are
indicated by "m/f."

Table A1. Birds

Natal Natal
dispersal dispersal
Body Obs
Species median maximum 1 2 3 n Source
mass (kg) type
distance distance
(km) (km)

Phalacrocorax 2.03 m
S ... 150.0 m/f C G M ... Hobson (1997)
pelagicus 1.7 f

11.80 m
Cygnus olor S ... 64.0 m/f H T R ... Ciaranca et al. (1997)
9.67 f

Cygnus 11.40 m
S ... 128.0 m/f O T M ... Mitchell (1994)
buccinator 10.30 f

Branta 3.814 m 2.0 m 28.4 m 19 m

D H T R Lessells (1985)
canadensis 3.314 f 1.5 f 11.2 f 20 f

0.681 m Hepp and Bellrose

Aix sponsa S ... 3.8 m/f O N M ...
0.685 f (1995)

Bucephala 1.00 m Dow and Fredga

D 0.75 f 6.0 f O N M 138 f
clangula 0.80 f (1983)

Bucephala
0.334 f S ... 4.5 f C T M ... Gauthier (1993)
albeola

Lophodytes 0.68 m
S ... 5.6 m/f C T M 10 f Dugger et al. (1994)
cucullatus 0.54 f

Eudocimus 1.04 m Kushlan and Bildstein

S ... 100.0 m/f C G M ...
albus 0.76 f (1992)

Elanus leucurus 0.325 m/f S ... 160.0 m/f C N R ... Dunk (1995)
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Marcström and
Accipiter 0.912 m 32 m 87 m Kenward (1981),
D 1150 m/f C T M
gentilis 1.137 f 17 f 66 f Mueller and Berger
(1967)

Accipiter 0.349 m 9.2 m Rosenfield and

D 35.2 m C T M 6m
cooperi 0.529 f 14.4 f Bielefeldt (1992)

0.150 m 9.0 m 108.0 m 99 m Newton and Marquiss

Accipiter nisus D C T R
0.280 f 18.5 f 166.0 f 65 f (1983)

Parabuteo 0.69 m
S ... 160.0 m/f C N R ... Bednarz (1995)
unicinctus 0.99 f

0.475 m
Buteo lineatus S ... 24.0 m/f C T M 4 m/f Crocoll (1994)
0.643 f

Buteo Preston and Beane

1.069 f S ... 320.0 f C T M 46 m/f
swainsoni (1993)

Aquila
3.0 m/f D 100.0 m/f 430.0 m/f C T R 24 m/f Ferrer (1993)
adalberti

Mearns and Newton

Falco 0.611 m 58.0 m 357.0 m
D C T R 11 m (1982), James et al.
peregrinus 0.952 f 83.0 f 324.0 f
(1989)

Falco 0.111 m 4.8 m 32.45 m Miller and Smallwood

D C T M 21 f
sparverius 0.120 f 5.06 f 38.79 f (1997)

5.7 m
Bonasa bonasia ... S ... H N R 9 m/f Fang and Sun (1997)
4.8 f

Dendragapus 0.492 m 0.6 m 6.0 m Hines (1986),

D H T M 88 m
canadensis 0.456 f 5.0 f 6.0 f Schroeder (1986)

Beaudette and Keppie

Dendragapus 1.188 m 0.9 m 2.6 m 24 m
D H T R (1992), Jamieson and
obscurus 0.891 f 1.4 f 11.0 f 42 f
Zwickel (1983)

Tympanuchus 0.999 m Bowman and Robel

D 1.0 m/f 10.8 m/f H N M 24 m/f
cupido 0.772 f (1977)

Meleagris 7.40 m
S ... 48.0 f O N R ... Eaton (1992)
gallopavo 4.22 f

Lagopus 0.359 m 1.0 m 7.5 m 258 m Giesen and Braun

D H T R
leucurus 0.516 f 4.0 f 29.0 f 68 f (1993)

Lagopus 0.601 m 1.0 m 4.0 m 60 m Martin and Hannon

D H T R
lagopus 0.516 f 2.9 f 7.5 f 17 f (1987)

Actitis 0.477 m
S ... 146.7 m/f C T P 5 m/f Oring et al. (1997)
macularia 0.394 f

Numenius 0.36 m Skeel and Mallory

S ... 1.63 m O N M ...
phaeopus 0.40 f (1996)

Recurvirostra 5m Robinson and Oring

0.32 m/f S ... 19.4 f O G M
americana 3f (1997)
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Charadrius 596.0 m
0.552 m/f D 11.8 m/f C T M 34 m/f Haig and Oring (1988)
melodus 146 f

Larus 0.57 m 2427

D 98.0 m/f 700 m/f O G M Gabrey (1996)
delawarensis 0.47 f m/f

Butler et al. (1980),

Larus 581
1.010 m/f D 100.0 m/f 400.0 m/f C G R Belant and Dolbeer
glaucescens m/f
(1993)

1836
Larus fuscus 0.71 m/f D 42 m/f 202 m/f C G R Paradis et al. (1998)
m/f

Columba 751
0.49m/f D 4 m/f 150 m/f O T R Paradis et al. (1998)
palumbus m/f

Zenaida 0.123 m 4.8 m Tomlinson et al.

S ... H N M ...
macroura 0.115 f 4.8 f (1960)

Belthoff and Ritchison

0.167 m (1989), VanCamp and
Otus asio D 10.7 m/f 158.5 m/f C N R 17 m/f
0.194 f Henny (1975),
Gehlbach (1986)

0.454 m
Strix aluco D 4.0 m/f 22.4 m/f C T R 9 m/f Southern (1970)
0.478 f

0.79 m Bull and Duncan

Strix nebulosa S ... 50.0 m/f C N R 21 m/f
1.16 f (1993)

Gutiérrez et al. (1985),

Strix 0.582 m
D 29.0 m/f 87.4 m/f C T R 11 m/f Miller and Meslow
occidentalis 0.637 f
(1985)

Speotyto
0.150 m/f S ... 30.0 m/f C G M ... Haug et al. (1993)
cunicularia

Wallin and Andersson

(1981), Löfgren et al.
23 m/f
Aegolius 0.101 m 12.8 m 67.5 m (1986), Korpimäki
D C N R 13 m
funereus 0.167 f 56.0 f 300.0 f (1987), Korpimäki
37 f
and Lagerström
(1988)

Bubo 1.154 m 111 Adamcik and Keith

S ... 1305.0 m/f C T R
virginianus 1.555 f m/f (1978)

Picoides
0.044 m/f S ... 90.0 m/f O T M ... Jackson (1994)
borealis

Dryocopus 0.31 m Bull and Jackson

S ... 32.0 m/f O T R 8 m/f
pileatus 0.27 f (1995)

Hirundo fulva 0.020 m/f S ... 30.0 m/f C G M ... West (1995)

Allen and Nice

Hirundo rustica 0.19 m/f D 6.4 m/f 8.1 m/f C G R 7 m/f
(1952), Shields (1982)

Iridoprocne
0.02 m/f D 1.0 m/f 6.4 m/f O N M 41 m/f Chapman (1955)
bicolor
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3046
Riparia riparia 0.015 m/f D 6.0 m/f 144.0 m/f C N M Mead (1979)
m/f

Progne subis 0.049 m/f D 40 m/f 336.0 m/f O G M 46 m/f Allen and Nice (1952)

Delichon
0.020 m/f D 0.03 m/f 4.3 m/f C N M 49 m/f Rheinwald (1975)
urbica

Aphelocoma 0.7 m 1.1 m 38 m Woolfenden and

0.080 m/f D H N R
coerulescens 0.3 f 1.8 f 32 f Fitzpatrick (1978)

Malurus 0.20 m Russell and Rowley

0.010 m/f D 2.40 m/f C G R 94 f
splendens 0.20 f (1993)

Gymnorhinus 1064 Marzluff and Balda

0.103 m/f S ... 640.0 m/f O N R
cyanocephalus m/f (1989)

Sayornis 0.02 m 34.4 m 33 m

S ... C T R Wolf (1997)
nigricans 0.018 f 20.0 f 30 f

Perisoreus 0.0 m
0.071 m/f D 11.3 m/f O T R 15 m/f Strickland (1991)
canadensis 2.8 f

0.189 m 0.35 m 1.3 m

Pica pica D O N R ... Eden (1987)
0.166 f 0.5 f 0.8 f

Parus 58 m Weise and Meyer

0.011 m/f D 1.1 f 11.2 f O N R
atricapillus 45 f (1979)

Berndt and Sternberg

(1968), Greenwood et
0.02 m 0.6 m 3.3 m 92 m
Parus major D O T R al. (1979), Weise and
0.018 f 0.9 f 3.3 f 98 f
Meyer (1979), Nilsson
(1989)

1.1 m 4.6 m 65 m
Parus palustris 0.014 m/f D O T R Nilsson (1989)
2.6 f 7.3 f 53 f

Parus Berndt and Sternberg

0.011 m/f D 0.7 m/f 470.0 m/f O T R 72 f
caeruleus (1968)

Berndt and Sternberg

1.1 m 29 m
Sitta europea 0.022 m/f D ... O T M (1968), Mattysen and
0.8 f 26 f
Schmidt (1987)

Vireo griseus 0.011 m/f S ... 20.0 m/f C T M ... Hopp et al. (1995)

Lanius Collister and De Smet

0.047 m/f D 7.3 m/f 79.7 m/f O T M 69 m/f
ludovicianus (1997)

Allen and Nice

74 m
Turdus merula 0.095 m/f D 5.0 m/f 355.0 m/f O T R (1952), Greenwood
50 f
and Harvey (1977)

2204
Turdus merula 0.095 m/f D 4.0 m/f 355 m/f O T R Paradis et al. (1998)
m/f

Toxostoma
0.079 m/f S ... 30.0 m/f C T R ... Tweit (1996)
curvirostre

Hylocichla 0.044 m 680

S ... 4.68 m/f C T M Anders et al. (1998)
mustelina 0.050 f m/f
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Acrocephalus
0.012 m/f D 51 m/f 271 m/f C N M 78 m/f Paradis et al. (1998)
scirpaceus

Motacilla alba 0.24 m/f S ... 100.0 m/f C N M ... Badyaev et al. (1996)

Seiurus 0.020 m
S ... 4.0 m/f C T M ... Robinson (1995)
motacilla 0.021 f

Passerculus 0.248 f 1.4 f 65 m Wheelwright and

0.019 m/f D H N M
sandwichensis 0.202 m 1.6 m 76 f Mauck (1998)

Nice (1937), Johnston

Melospiza 0.20 m 13.2 m 36 m/f (1956), Halliburton
0.021 m/f D H T R
melodia 0.20 f 1.3 f 34 m/f and Mewaldt (1976),
Arcese (1989)

Zonotrichia 0.3 m 138 m Baker and Mewaldt

0.294 m/f D 2.6 m/f O T M
leucophrys 0.4 f 173 f (1978)

Passerina
0.108 m/f S ... 470.0 m/f O N M ... Payne (1991)
cyanea

Carduelis 7.0 m 150.0 m 15 m Greenwood and

0.03 m/f D H T R
chloris 3.0 f 37.5 f 11 f Harvey (1977)

Ficedula 473 Berndt and Sternberg

0.015 m/f D 0.9 m/f 75.0 m/f O T M
hypoleuca m/f (1968), Pärt (1990)

Quiscalus 0.21 m 35.0 m

S ... O G R ... Post et al. (1996)
major 0.12 f 7.0 f

0.05 m
Molothrus ater S ... 40.0 m/f O N M 15 m/f Lowther (1993)
0.04 f

Table A2. Mammals

Natal Natal
Body
Obs dispersal dispersal
Species mass 1 2 n Source
type median maximum
(kg)
distance (km) distance (km)

Didelphis 2.839 m 4.3 m

S ... O T 8 VanDruff (1969)
virginianus 1.99 f 5.152 f

Phascogale 0.199 m Soderquist and Lill

S ... 6.8 m C T ...
tapotafa 0.145 f (1995)

Trichosurus
2.93 m/f D 5.4 m 12.8 m C T 13 m Cowan et al. (1996)
vulpecula

Hanski et al. (1991),

29 m
Sorex araneus 0.004 m/f S ... 0.869 m/f C N Peltonen and Hanski
21 f
(1991)

Scapanus 0.722 m 44
0.148 m/f S ... O G Carraway et al. (1993)
townsendii 0.856 f m/f

Ursus 130.0 m 225.0 m 19 m

S ... O T Rogers (1987)
americanus 78.9 f 28.8 f 15 f
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134.0 m 35 m Glenn and Miller

Ursus arctos 331.0 m/f S ... O T
82.0 f 26 f (1980)

222
5.5 m
Ursus arctos 331.0 m/f D ... O T m Pearson (1972)
9.5 f
343 f

8.1 m 34 Priewert (1951),

Procyon lotor S ... 265.5 m/f O T
7.5 f m/f Lynch (1967)

Martes 15
1.04 m/f S ... 61.0 m/f C T Lofroth (1993)
americana m/f

Martes 3.70 m 23.0 m 35

S ... C T Arthur et al. (1993)
pennanti 2.10 f 22.6 f m/f

Mustela 0.080 m 5.6 m 36 m

S ... C T Erlinge (1977)
erminea 0.054 f 1.0 f 29 f

1.225 m 45.1 m 6m
Mustela vison S ... C T Mitchell (1961)
1.112 f 45.0 f 2f

Lutra 39 Melquist and

9.07 m S ... 42.0 m C T
canadensis m/f Hornocker (1983)

Pasitschniak-Arts and
Gulo gulo 14.4 m/f S ... 300 m/f C T ...
Lariviere (1995)

7.80 m 110.0 m 1m
Taxidea taxus S ... C T Lindzey (1978)
6.15 f 52.0 f 1f

11.6 m 8.3 m
Meles meles S ... O G ... Cheesman et al. (1988)
10.1 f 7.8 f

Mephitis 3.176 m 10.1 m 1m

S ... O T Bjorge et al. (1981)
mephitis 2.642 f 21.7 f 4f

Helogale 0.25 m 44 m
0.350 m/f D ... C T Rood (1987)
parvula 0.75 f 22 f

176.0 m Andrews and Boggess

Canis latrans 14.06 m/f S ... C N 125
232.2 f (1978)

16.6 m 7m
Canis latrans 14.06 m/f D ... C N Bowen (1982)
42.2 f 5f

42.54 m 432.0 m 5m Berg and Kuehn

Canis lupus S ... C T
40.22 f 79 f 2f (1982), Mech (1987)

42.54 m 125.0 m 9m
Canis lupus D ... C T Ballard et al. (1987)
40.22 f 125.0 f 5f

42.54 m 225.0 m 40 m
Canis lupus D ... C T Gese and Mech (1991)
40.22 f 75.0 f 29 f

394.5 m 1m Ables (1965), Allen

Vulpes vulpes 5.20 m/f S ... C T
302.0 f 72 f and Sargent (1993)

8.6 m 31 m
Vulpes vulpes 5.20 m/f D ... C T Jensen (1973)
4.9 f 31 f
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298
24.2 m
Vulpes vulpes 5.20 m/f D ... C T m Storm et al. (1976)
8.1 f
207 f

171
8.1 m
Vulpes vulpes 5.20 m/f D ... C T m Phillips et al. (1972)
8.1 f
124 f

12.0 m 16 m
Vulpes vulpes 5.20 m/f D ... C T Zimen (1984)
12.4 f 4f

Urocyon
4.1 m
cinereoargente S ... 83.68 f C T ... Sullivan (1956)
3.9 f
us

Logan et al. (1986),

61.6 m 274.0 m 33 m
Felis concolor S ... C T Ross and Jalotsky
44.5 f 155.0 f 13 f
(1992)

Lynx lynx 10.149 f S ... 9.7 f C T 1 Saunders (1963)

Robinson and Grant

Lynx rufus 8.2 f S ... 56.0 f C T 10
(1958)

Lariviere and Walton

Lynx rufus 9.6 m S ... 182 m C T ...
(1997)

Castor 19
24.3 f S 2.1 m/f 40.6 m/f H T Leege (1963)
canadensis m/f

Armitage (1974),
Marmota
5.448 m S ... 1.4 m H G 2 Armitage and
flaviventris
Downhower (1974)

Marmota 3.511 m 0.685 m 46

S ... H N Swihart (1992)
monax 3.486 f 0.768 f m/f

Spermophilus 0.05 m 0.975 m 26 m Mitchener and

0.405 m/f D H G
richardsonii 0.05 f 0.525 f 26 f Mitchener (1977)

Spermophilus 1.44 m 56 m Evans and Holdenried

0.475 m/f S ... H G
beecheyi 1.312 f 25 f (1943)

Spermophilus 25 m
0.400 m/f S ... 0.328 m/f H G Holekamp (1984)
beldingi 2f

Spermophilus 0.515 m 65 m Allred and Beck

0.192 m/f S ... H N
leucopus 0.321 f 83 f (1963)

Spermophilus
46 m
tridecemliniatu 0.140m S ... 0.239 m H T Rongstad (1965)
45 f
s

Sciurus niger 0.790 m/f S ... 3.37 m/f H T 2 Fitch (1958)

Tamiasciurus 0.204 m 0.485 m 0.60 m

D H T 8 m/f Sun (1997)
hudsonicus 0.199 f 0.24 f 0.60 f

Thomomys 2m Daly and Patton

0.150 m/f S ... 0.30 m/f C T
bottae 1f (1990)
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Perognathus 28 m Allred and Beck

0.023 m S ... 0.853 H T
formosa 18 f (1963)

Dipodomys 0.044 m 0.13 m 0.25 m 13 m

D H T Jones (1989)
merriami 0.0426 f 0.03 f 0.158 f 13 f

159
Dipodomys 0.411 m Allred and Beck
0.065 m/f S ... H T m
microps 0.434 f (1963)
172 f

Dipodomys 0.9 m
0.115 m/f S ... H T ... Jones (1987)
spectabilis 2.0 f

Dipodomys 0.03 m 82 m Waser and Elliott

0.115 m/f D ... H T
spectabilis 0.03 f 84 f (1991)

Dipodomys 0.03 m 22 m
0.115 m/f D ... H T Jones (1987)
spectabilis 0.03 f 18 f

Dipodomys 0.087 m 13 m
0.064 m/f D ... H T Price et al. (1994)
stephensi 0.045 f 18 f

Neotoma 0.42 m
S ... 2.2 m H T ... Smith (1997)
cinerea 0.30 f

Clethrionomys 58 Dickman and

0.023 m/f S ... 0.40 m/f H T
glareolus m/f Doncaster (1989)

Onychomys 0.329 m 1m Allred and Beck

26.8 m/f S ... H T
torridus 0.325 f 1f (1963)

Microtus 0.03 m 0.159 m 39 m

0.023 m/f D H T Sandell et al. (1990)
agrestis 0.03 f 0.193 f 19 f

Microtus Boyce and Boyce

0.022 f D 0.03 f 0.2 f H N 31 f
arvalis (1988)

Microtus 0.028 m 0.136 m 79 m

S ... H T McGuire et al. (1993)
ochrogaster 0.033 f 0.127 f 78 f

Microtus
0.032 m/f D 0.75 m 1.0 m H T 5m Steen (1994)
oeconomus

Microtus
0.040 m/f S ... 0.245 m/f H T 1 Mihok et al. (1988)
pennsylvanicus

Microtus 0.008 m 0.068 m 52 m

0.032 m/f D H T Lambin (1994)
townsendi 0.003 f 0.054 f 300 f

Microtus Wolff and Lidicker

135 m/f S ... 0.3 m/f H G ...
xanthognathus (1980)

Peromyscus 0.06 m 0.45 m 27 m

0.018 m/f D H T Ribble (1992)
californicus 0.15 f 0.791 f 24 f

Peromyscus 0.228 m 1m Allred and Beck

0.008 m/f S ... H T
longicaudus 0.235 f 1f (1963)

Peromyscus 0.05 m 0.883 m 71 m Dice and Howard

0.015 m/f D O T
maniculatus 0.15 f 1.005 f 64 f (1951)
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Apodemus 261 Dickman and

0.020 m/f S ... 0.5 m/f H N
sylvaticus m/f Doncaster (1989)

Ondatra
1.048 m/f S ... 3.37 m/f H T ... Fitch (1958)
zibethicus

Ochotona 15 Peacock and Smith

... D 0.09 m/f 0.396 m/f H G
princeps m/f (1997)

Lepus
1.43 m/f S ... 20.1 m/f H T 3 O’Farrell (1965)
americanus

Broekhuisen and
Lepus europea 4.50 m/f S ... 9.0 m/f H T 11
Maaskamp (1982)

99 Angerbjörn and Flux

Lepus timidus 2.750 m/f S 3.0 m/f 200 m/f H N
m/f (1995)

Lepus
2.2 m/f S ... 45 m/f H N ... Best (1996)
californicus

Sylvilagus 148
0.841 m/f S ... 0.352 m/f H N Shields (1960)
bachmani m/f

Sylvilagus 1.10 m 3.9 m 65 Chapman and

S ... H T
floridanus 1.20 f 2.3 f m/f Trethewey (1972)

Odocoileus
64.638 4.0 m 15.2 m 26 m Bunnell and Harestad
hemionus D H G
m/f 2.0 f 12.2 f 14 f (1983)
columbianus

Odocoileus
82.5 m 7.34 m 65 m
hemionus S ... H G Robinette (1966)
51.75 f 8.22 f 33 f
hemionus

Odocoileus Nelson and Mech

86.41 f S ... 11.74 f H G 7f
virginianus (1992)

Odocoileus 6.0 m 26 m
86.41 f D ... H G Nelson (1993)
virginianus 24.0 f 4f

481.83
Alces alces S ... 118.0 H T 87 Pullianen (1974)
m/f

Cervus 315 m
S ... 18.5 m/f H T 5 m/f Brazda (1953)
canadensis 225 f

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